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Equine Reproductive
Physiology, Breeding
and Stud
Management
5th Edition
Equine Reproductive
Physiology, Breeding
and Stud
Management
5th Edition
CABI
CABI is a trading name of CAB International
CABI CABI
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E-mail: [email protected]
Website: www.cabi.org
© Mina C.G. Davies Morel 2021. All rights reserved. No part of this publication may be reproduced in any form or by any means,
electronically, mechanically, by photocopying, recording or otherwise, without the prior permission of the copyright owners.
A catalogue record for this book is available from the British Library, London, UK.
Names: Davies Morel, Mina C. G., author. Title: Equine reproductive physiology, breeding and stud management / Mina C.G. Davies
Morel.
Description: Fifth edition. I Boston : CAB International, 2021. I Includes bibliographical references and index. I Summary: "Equine
Reproductive Physiology Breeding and Stud Management, 5th Edition provides a thorough grounding in equine reproductive anatomy
and physiology for equine, animal and veterinary science students"-- Provided by publisher.
Identifiers: LCCN 2020018102 (print) I LCCN 2020018103 (ebook) I ISBN 9781789242249 (paperback) I ISBN 9781789242232
(hardback) I ISBN 9781789242256 (ebook) I ISBN 9781789242263 (epub)
Subjects: LCSH: Horses--Reproduction. I Horses--Breeding.
Classification: LCC SF768.2.H67 D39 2020 (print) I LCC SF768.2.H67 (ebook) I DDC 636.l--dc23
LC record available at https://lccn.loc.gov/2020018102
LC ebook record available at https://lccn.loc.gov/2020018103
Glossary 399
Bibliography 403
Index 507
Videos for Equine Repro Phys
Breeding and Stud Management
Breeding Soundness evaluation in the mare including ultrasound examination, vaginascopy, swabbing and uterine biopsy
• https:/ /www.youtube.com/watch?v=srZhtfLFVFg
Excerpts from a longer video describing the process of ultrasound examination in the mare
• https://www.youtube.com/watch?v=Q-xR6ez 1 h7I&t=7s
[!].
AI, AV preparation, semen collection, semen evaluation and insemination into the mare
• https://www.youtube.com/watch?v=ohei 1 Kl s7As
Semen evaluation
• https://www.youtube.com/watch?v=NCAjYl IOHTc&index=3&list=PLBPI2E05ryn-XJOcNBH
WZFT-FyKsoVslf
Videos for Equine Repro Phys Breeding and Stud Management
Flushing the mare, identifying the embryo and transferring the embryo into the recipient mare
• https://www.youtube.com/watch?v=78-XVxHqruA
Videos for Equine Repro Phys Breeding and Stud Management
Section A considers the biology of breeding the mare, the is controlled in the mare. This knowledge will then en
anatomy of the mare, the processes involved in ova pro able you to understand the following sections, which
duction (folliculogenesis and oogenesis), fertilization, apply this knowledge to breeding practice.
pregnancy and parturition, and how reproductive activity
Mare Reproductive Anatomy
© CAB International 2021. Equine Reproductive Physiology, Breeding and Stud Management,
5th Edition. (M. Davies Morel.)
Chapter 1
I
,I
Fig. 1.3. A lateral (side) view of the mare's reproductive tract.
Fig. 1.4. The vulval area of the mare: in this instance, the con
formation of the perinea! area is poor, with the anus sunken
cranially, opening up the vulva to faecal contamination. (Photo
courtesy of Ms Ria McLean.)
-----
Vestibular (vaginal) seal Vulval seal
Fig. 1.6. The seals of the mare's reproductive tract during dioestrus.
(A) (B)
(C)
Fig. 1.7. The effect of conformation on the competence of the vulval, vestibular and cervical seals in the mare: (A) a low ischium
(pelvic Aoor) results in an incompetent vestibular seal: in this case, the vulval seal is still competent, so infection risk is limited; (B) a low
ischium results in an incompetent vestibular seal: in this case, the vulval seal is also incompetent, so infection risk is increased; and
(C) an incompetent vestibular and vulval seal plus a sloping perinea! area result in a significant infection risk, especially from faecal
contamination.
Mare Reproductive Anatomy
inserted through the vulva into the vagina and allowed tone, tends to draw air into the tract, especially when
to rest horizontally on the vagina floor, the amount of the mare is moving or coughs (Hemberg et al., 2005).
vulva lying below this tube should be approximately Pascoe (1979) suggested that mares should be allocated
80% in well-conformed mares. This technique is illus a Caslick index derived by multiplying the angle of in
trated diagrammatically in Fig. 8.7. clination of the vulva with the distance from the is
If the ischium of the pelvis is too low, the vulva chium to the dorsal commissure. This index can then be
tends to fall towards the horizontal plane as seen in Figs used to classify mares into three types and so predict the
1.4 and 1.7. This opens up the vulva to contamination likely occurrence of endometritis (Fig. 1.9).
by faeces, increasing the risk of uterine infection due to The effect of poor conformation of the perineum
pneumovagina. Additionally, a low pelvis causes the area may be alleviated by a Caslick's vulvoplasty oper
vagina to slope inwards, preventing the natural drain ation, developed by Dr Caslick in 1937 (Caslick, 1937).
age of urine at urination leading to urovagina, which The lips on either side of the upper vulva are cut, and
further increases the risk of uterine infection. Lastly, the two sides are then sutured together. The two raw
negative abdominal pressure, typical of Thoroughbred edges then heal together as in the healing of an open
type, multiparous mares with poor abdominal muscle wound, and hence seal the upper part of the vulva and
prevent the passage of faeces into the vagina. The hole
left at the ventral commissure is adequate for urination
(Fig. 1.10). Short and long Caslick vulvoplasties may be
performed, depending on the severity of the perinea!
mal-conformation and hence on the length of vulva
that requires to be sutured.
The chance that a mare requiring a Caslick's oper
ation will pass on the trait for poor perinea! conform
ation to her offspring is reasonably high. This, coupled
with the fact that the operation site has to be cut to
allow natural mating and foaling, casts doubt on
whether such mares should be bred. It becomes increas
Fig. 1.8. The effect of oestrus on the competence of the vulval, ingly hard to perform a Caslick's operation on mares
vestibular and cervical seals in the mare: oestrus causes a relax that have been repeatedly cut and resutured, as the lips
ation of the seals and, therefore, an increase in infection risk. of the vulva become progressively more fibrous and
--------------= -
(A) (B) (C)
�
---- r
Dorsal
commissure
-------------- ----r
40%
Ventral
_______ l
Fig. 1.9. A lateral view of the relationship between the anus, vulva and ischium, indicating: (A) type I mare with good conformation,
Caslick index< 50 (b= 2-3 cm, a< 10 °): no Caslick required; (B) type II mare with poor conformation, predisposing to type III in later
life, Caslick index 50-100 (b = 6-7 cm, a= 10-15 °): no immediate need for a Caslick but likely in later life; and (C) type III mare with
very poor conformation, including vulva lips in a horizontal plane, Caslick index> 150 (b = 5-9 cm, a � 30 °): Caslick required immedi
ately, significant chance of endometritis and a reduction in reproductive success.
Chapter 1
Fig. 1.10. A Caslick operation in the mare showing (A) anaesthetizing the vulval lips; (B) cutting the vulval lips; (C) suturing together
the vulval lips; and (D) the finished job.
therefore difficult to suture. In such cases, a procedure Perinea! mal-conformation, and hence pneumova
termed a Pouret operation may be performed (Pouret, gina and urovagina, is particularly prevalent in Thor
1982). This is a more major operation and involves the oughbred mares. The condition tends to be exacerbated
surgical realignment of the anus and vulva (Knottenbelt in mares with a low body condition score; multiparous,
and Pascoe, 2003). Other alternatives have been sug aged mares; those in fit athletic condition; and those
gested, modified Pouret (Papa et al., 2014) and vulva! with flat croup and/or elevated tail set. Its continued
flap (Inoue and Sekiguchi, 2017), but are yet to be existence is in no small part due to the selection of hors
widely adopted. es for athletic performance rather than reproductive
Mare Reproductive Anatomy
competence (LeBlanc, 1991; Easley, 1993). Recent work protective mucus layer produced by mucus secretory
(McLean, 2014) indicated that up to 50% of Thor cells. The exact composition of vaginal secretion is con
oughbred mares in the UK have had a Caslick operation. trolled by the cyclical hormonal changes of the mare's
The oestrous cycle also has an effect on the compe reproductive cycle; see Chapter 2.
tence of the three seals. Further details of the effect of
the oestrous cycle on the reproductive tract are given in 1.5. The Cervix
Chapter 2. In summary, oestrus (period of sexual recep The cervix lies at the entrance to the uterus and is a re
tivity) results in the slackening of all three seals, due to markably versatile structure, normally providing a
a relaxation of the muscles associated with the repro tight, thick-walled sphincter, hence acting as the final
ductive tract, especially the cervix (Fig. 1.8). This allows protector of the system, but is also able to dilate vastly
intromission at covering but also decreases the compe to accommodate the passage of the fetus at parturition
tence of the reproductive tract seals and so increases the (Figs 1.12 and 1.13). The walls of the cervix form a ser
chance of bacterial invasion. In part, this is compen ies of folds or crypts and are highly muscular with col
sated for by elevated oestradiol levels characteristic of lagenous connective tissue cores and lined by folded
oestrus, which enhance the mare's immunological re columnar epithelium containing mucus-secreting cells.
sponse, so reducing the chance of uterine infection, These crypts are continual with the uterine endomet
despite the increased chance of bacterial invasion. rium folds and enable the significant expansion re
quired at parturition (Ginther, 1992; Kainer, 2011)
1.4. The Vagina (Fig. 1.14). In the sexually inactive, dioestrous state, the
The vagina of the mare is on average 18-23 cm long cervix is tightly contracted, white in colour and meas
and 10-15 cm in diameter. In the well-conformed mare ures on average 6-8 cm long and 4-5 cm in diameter;
the floor of the vagina should rest upon the ischium of
the pelvis, and the walls are normally collapsed and ap
posed, forming the vestibular seal. The hymen, if pre
sent, is also associated with this seal and divides the
vagina into anterior (cranial, nearest the mare's head)
and posterior (caudal, nearest the mare's tail) sections.
In some texts the posterior vagina is referred to as the
vestibule. The urethra, from the bladder, opens just cau
dal to the hymen. The walls of the vagina are muscular
and include the constrictor vestibule muscle. The pos
terior vagina is lined by stratified squamous epithelium
which accommodates abrasion at copulation whereas Fig. 1.11. The internal surface of the mare·s vagina illustrating
the anterior vagina is lined by columnar epithelium. In from the left the vulva; the posterior vagina lined by stratified
addition both the posterior and anterior vagina are squamous epithelium; the transverse fold (position of the
lined by mucus-secreting cells. The muscle layer pro hymen); the anterior vagina lined by columnar epithelium; the
vides elasticity and its dorsal incompleteness allows the cervix.
major stretching required at parturition (Fig. 1.11).
The vagina acts as the second protector and cleaner
of the system. It is largely aglandular (does not contain
secretory glands) but contains acidic to neutral secre
tions, originating from the mucus-secreting cells, the
cervix and small vestibular glands situated in the poster
ior vagina. These acidic secretions are bacteriocidal (kill
bacteria); however, they are also spermicidal (kill
sperm), necessitating that sperm are deposited into the
top of the cervix/bottom of the uterus at mating, to
avoid the detrimental effect of the acidic conditions.
The acidic conditions also attack the epithelial cell lin Fig. 1.12. The dioestrous cervix is retracted, presenting a tight
ing of the vagina, but these cells are protected by the seal against entry into the uterus
Chapter 1
Serous layer
Myometrial layer
\._,.._.,._,,.,....---t--t--- Endometrium
Fig. 1.15. A cross section through the abdomen of the mare illustrating the two uterine horns and supporting broad ligaments.
Mare Reproductive Anatomy
downwards into the abdominal cavity. The uterus of the endometrial glands and ducts (Figs 1.16 and 1.17). The
mare is termed a simplex bipartitus, due to the rela submucosa can be further divided into the compact
tively large size of the uterine body compared to the layer (stratum compactum), nearest the epithelium,
uterine horns (60:40 split). This differs from that in and the spongy layer (stratum spongiosum), nearest the
other farm livestock, where the uterine horns are more myometrium. Collagenous connective tissue cores sup
predominant. The lack of a septum dividing the uterine port these endometrial folds. The activity and, there
body is also notable (Hafez and Hafez, 2000; Frandson fore, appearance of the endometrial glands and the
et al., 2009). In situ the uterine walls are flaccid and epithelial cells are dependent on the cyclical hormonal
intermingle with the intestine, the only lumen present changes associated with the oestrous cycle. It is the
being very small and that formed between the endo endometrium that is responsible for supporting the
metrial folds. early conceptus and for placental attachment and devel
The uterine wall (Fig. 1.16), in common with the opment (Ginther, 1992, 1995; Senich, 1998; Kainer,
rest of the tract but most prominent here, consists of 2011). Causey (2007) also suggested that within the
three layers: the perimetrium (an outer serosa layer) uterine epithelium are mucus-secreting and ciliated
continuous with the broad ligaments; the myometrium cells that help eliminate bacteria, providing an add
(central muscular layer); and the inner endometrium. itional defence against uterine bacterial invasion.
The central myometrial layer is particularly evident in
the uterus where clearly defined outer longitudinal 1.7. The Utero-tubular Junction
muscle fibres, a central vascular layer and inner circular The utero-tubular junction is a constriction or sphinc
muscle fibres can be seen. It is this central myometrial ter formed by a high concentration of muscle cells ori
layer that allows the elasticity for expansion of the ginating from the circular myometrium of the Fallopian
uterus during pregnancy as well as providing the force tube. The junction, which appears as a papilla in the
for parturition. The inner endometrium is arranged in endometrium, provides an abrupt connection between
12-15 longitudinal folds continuous with the folds of the end of the uterine horns and the Fallopian tubes
the cervix (Figs 1.2 and 1.14) and comprises the outer (Fig. 1.18). Fertilization takes place in the Fallopian
epithelial cells (epithelium) and inner submucosa of tubes, and the utero-tubular junction selectively allows
endometrial connective or stroma tissue with its associated only fertilized ova to pass through and on to the uterus
Epithelium
Endometrial glands
Strama
Endometrium
Myometrium
Perimetrium
Fig. 1.16. A cross section through the uterine wall. Apart from the presence of endometrial glands, which are present in just the
uterus, this cross section is the same throughout the whole of the reproductive tract.
Chapter 1
Fig. 1.17. A uterine biopsy illustrating a cross section through the uterine wall, illustrating from the top left: the outer epithelial cells
of the endometrium; the endometrium stratum compactum, with a few endometrial glands; the endometrium stratum spongiosum,
with a high concentration of endometrial glands; the circular muscle cells of the myometrium; and finally, in the very bottom right
hand corner, the beginnings of the longitudinal muscle cells of the myometrium.
which pushes the fertilized ova towards the uterotubu 6-8 cm in length and 3-4 cm in width; they are also softer
lar junction. The ampulla of each Fallopian tube ends in to the touch owing to the development of fluid-filled fol
the infundibulum, a funnel-like opening close to the licles (Fig. 1.20). Older, multiparous mares tend to show
ovary (Kainer, 2011). larger ovaries which can be up to 10 cm in length.
The infundibulum in the mare is closely associated The mare's ovaries are bean shaped with the convex
with a specific part of the ovary, termed the ovulation outer surface or border of the ovary attached to the
fossa, which is unique to the mare and is the only site of mesovarian section of the broad ligaments (Figs 1.20
ova release; in other mammals ovulation may occur and 1.21), which is the entry point for blood and nerve
over the whole surface of the ovary and so the infun supply; the concave inner surface is free from attach
dibulum encapsulates the ovary. The infundibulum is, ment and is the location of the ovulation fossa. The
therefore, relatively hard to distinguish in the mare, not whole ovary is contained within a thick protective
being so evident as a funnel-shaped structure surround layer, the tunica albuginea, except for the ovulation
ing the whole ovary. Like the ampulla, the infundibu fossa. The tissue of the ovary in the mare is arranged as
lum is lined by cilia, which again attract and catch the the inner cortex (active gametogenic and steriodogenic
ova guiding them towards the entrance of the Fallopian tissue) and the outer medulla (supporting tissue). Ova
tubes (Ginther, 1992; Kainer, 2011). release at ovulation occurs only through the ovulation
fossa, and all follicular and corpus luteum (CL) devel
1.9. The Ovaries opment occurs internally, within the cortex of the
The ovaries of the mare are both gametogenic (site of ovary (Fig 1.21). The mare differs in these aspects from
gamete (ova) production) and steroidogenic (site of hor other mammals, in which the medulla and cortex are
mone production) in function. They are evident as two reversed, ovulation occurring over the surface of the
bean-shaped structures normally situated ventrally to ovary and all follicular and CL development occurring
(below) the fourth and fifth lumbar vertebrae and sup on the outer borders. Rectal palpation, as a clinical aid
ported by the mesovarium part of the broad ligaments. to assess reproductive function in the mare, is not,
They make the total length of the reproductive tract in therefore, as easy to perform as it is in other farm live
the mare in the region of 50-60 cm. In the sexually in stock such as the cow. However, with the advent of
active stage, i.e. during the non-breeding season, the ultrasound, assessment of ovarian characteristics in the
mare's ovaries measure around 2-4 cm in length and 2-3 mare is now very accurate (see Section 10.2.2.2
cm in width and are hard to the touch owing to the ab Ginther, 1992, 1995; Senich, 1998; Hafez and Hafez,
sence of developing follicles. During the sexually active 2000; Kainer, 2011).
stage when the mare is in the breeding season, particu
larly during oestrus, they increase in size to around
Fig. 1.19. The convoluted Fallopian tube running through the Fig. 1.20. The ovaries of the mare. Note the difference in size
mesovarian section of the broad ligaments, from the ovary on between the ovary on the right (inactive) and the one on the
the left to the uterine horn on the right, illustrating the broader left (active). The concave surface (position of the ovulation
ampulla region of the Fallopian tube (on the left) and the more fossa) and the convex surface (the hilus, entry point for blood
wiry and narrower isthmus region to the right. and nerve supply) of the ovary are clearly seen.
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